A widespread neotropical species, ranging from the southern U. S. to subtropical southern South America and throughout the Antilles.
Etymology: The Charites, or Graces, are the personifications of charm and beauty in nature and in human life. They love all things beautiful and bestow talent upon mortals. Together with the Muses they serve as sources of inspiration in poetry and the arts. Originally, they were goddesses of fertility and nature, closely associated with the underworld and the Eleusinian mysteries (Charites).
Papilio charitonia L. is an objective junior synonym of Papilio charithonia L. The latter is on the Official List of Generic and Specific Names in Zoology (see Holthuis and Hemming 1956). See also Brower (1994). (NB: Linnaeus placed all butterflies he described in the genus Papilio. Thus the original name as described in 1767 is Papilio charithonia Linnaeus. Since 1802, this species has been assigned to the genus Heliconius Kluk.
There should not be a separate page for "Heliconius charitonia" in EOL.
- Brower, A.V.Z. 1994. The case of the missing H: Heliconius charithonia (L., 1767), not "Heliconius charitonia (L. 1767)". J. Lepid. Soc. 48: 166-168.
- Holthuis, L. B. and F. Hemming. 1956. Opinion 382: validation under the Plenary Powers of the generic name "Sicyonia" Milne Edwards (H.) 1830 (Class Crustacea, Order Decapoda) and action consequential thereon. pp. 43-58 in: Hemming, F. (ed.). Opinions and Declarations Rendered by the International Commission on Zoological Nomenclature 12(3).)
North American Ecology (US and Canada)
- Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
A widespread neotropical species, ranging from the southern U. S. to subtropical southern South America and throughout the Antilles.
This information is based an ongoing project dedicated to the inventory and dissemination of information on lepidopteran larvae, their host plants, and their parasitoids in a Costa Rican tropical wet forest and an Ecuadorian montane cloud forest.
N=10 rearings as of 2012, 9 eclosed and 1 died.
The Zebra Longwing (Heliconius charithonia) is a primarily neotropical butterfly with a distribution that extends north through the Florida peninsula in the U.S. (it is Florida's official state butterfly), sometimes ranging farther north as well. It has long, rounded wings and a slender body. It is pitch black with bold, yellow stripes and small red dots close to the body (Cech and Tudor 2005).
H. charithonia occurs from sea level to 1,800 m in edges and scrubs. Usually individuals fly erraticly and in the lowerstory. The males sit on female pupae a day before emergence, and mating occurs the next morning, before the female has completely eclosed. Adults roost at night in large groups lower than 2 m above ground in twigs or tendrils (Brown, 1981).
Host plant: H. charithonia larvae feed primarily on plants from the genus Passiflora, subgenus Granadilla, Tryphostemmatoides, and Plectostemma (Brown, 1981). In Costa Rica H. charithonia feeds on Tetrastylis lobata (Passifloraceae) (DeVries, 1997).
Heliconius charithonia on passion-vine in Mexico City, Mexico. © Maria Franco
Geographical distribution and named races
Heliconius charithonia is distributed from North America (Texas and Florida) to Venezuela and Peru, and also occurs on the Greater and Lesser Antilles. Although the species does not exhibit the dramatic geographical variation in wing patterns of some of its congeners, a number of names have been applied to various island populations.
H. charithonia charithonia (L., 1767)"America"(St. Thomas, Virgin Is., according to Comstock and Brown, 1950).
H. charithonia antiquus Lamas, 1988 (replacement name for H. charithonia punctata A. Hall, 1936 (preoccupied by h. cydno f. punctata Neustetter, 1907) (Antigua and St. Kitts)
H. charithonia bassleri Comstock & Brown, 1950 (Colombia)
H. charithonia churchi Comstock & Brown, 1950 (Haiti)
H. charithonia ramsdeni Comstock & Brown, 1950 (Cuba)
H. charithonia simulator Comstock & Brown, 1950 (Jamaica)
H. charithonia tuckeri Comstock Brown, 1950 (USA: Florida)
H. charithonia vasquezae Comstock & Brown, 1950 (Mexico: Campeche)
Early stages: Eggs are yellow or white and approximately 1.2 x 0.8 mm (h x w). Females usually place 1 to 5 eggs on growing shoots of the host plant. Mature larvae have a white body, with black spots or bands, black scoli and yellow and black or black and white head; length is around 1.2 cm. Caterpillars are gregarious in small numbers (Brown, 1981).
From left to right: Heliconius charithonia female laying eggs on her hostplant in Mexico City. © Maria Franco. Eggs, second, and third instar larva on hostplant in Gainesville, Florida. In Florida, H. charithonia larvae often completely defoliate their Passiflora hostplants. © Andrew V. Z. Brower. Last instar larva feeding on host plant and getting ready to pupate in Mexico City. © Maria Franco.
From left to right: Heliconius charithonia early pupa, pupa with butterfly ready to eclose, eclosing butterfly, and eclosed butterfly expanding and drying its wings. Mexico City, Mexico, November 2005 © Maria Franco.
Adult: Distinguished immediately by the zebra pattern, which gives it the common name of the "zebra" (DeVries, 1997).
Collected in Heredia Province, Costa Rica.
occurs (regularly, as a native taxon) in multiple nations
Regularity: Regularly occurring
Type of Residency: Year-round
Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) Texas to Florida and south to the Keys. Wanders as far as California, Great Basin, and the Great Plains; sometimes colonizes as far north as South Carolina. Also occurs in Mexico, the West Indies, and South America.
In the United States, the Heliconius charitonius is unmistakable. Over much of its Neotropical distribution, there are a dazzling array of other Heliconius species with which it might be confused, although the lack of any conspicuous colors other than black and yellow sets it apart from several otherwise similar species (Cook et al. 1976).
In Florida, the Zebra Longwing is found in subtropical hammocks (closed canopy forests, typically slightly higher in elevation than the surrounding area, that are dominated by a diverse assemblage of broad-leaved evergreen trees and shrubs, mostly of West Indian origin) and pine-oak woods, as well as in suburbs, parks, and even open fields if preferred flowers are available (Cech and Tudor 2005). Over must of its range to the south, this species is found in moist subtropical and tropical forests with sunlit openings (Opler and Krizek 1984).
Comments: Tropical lowland forests edges or openings; scrub. Larval hosts are in the genus Passiflora. Sometimes suburban habitats.
Non-Migrant: No. All populations of this species make significant seasonal migrations.
Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: No. No populations of this species make annual migrations of over 200 km.
Like many other Heliconius butterflies, H. charithonia feeds extensively on pollen, using its specially modified proboscis, as a source of amino acids and other nutrients (Gilbert 1972; Opler and Krizek 1984; Cech and Tudor 2005).
Passifloraceae: Passiflora lobata
Heliconius charithonia caterpillars feed on plants in the genus Passiflora (passion vines/passionflower/passionfruit). In the U.S., hosts include P. incarnata, P. lutea, P. suberosa, and P. multiflora (Cech and Tudor 2005).
Zebra Longwings are poisonous and distasteful to vertebrate predators (Opler and Krizek 1984; Cech and Tudor 2005).
Number of Occurrences
Note: For many non-migratory species, occurrences are roughly equivalent to populations.
Estimated Number of Occurrences: 81 to >300
10,000 to >1,000,000 individuals
Life History and Behavior
- Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
Male Zebra Longwings patrol for females and are attracted to female chrysalids, with whom they seek to mate at or near the time of emergence; after mating, the male deposits an "antiaphrodisiac" chemical onto the female's abdomen to deter matings with additional males (Cech and Tudor 2005). The typical flight pattern of the Zebra Longwing is slow and direct with very shallow, rapid wingbeats (Opler and Krizek 1984), although these butterflies are capable of faster flight when alarmed (Cech and Tudor 2005).
Each evening, Zebra Longwings gather to form communal sleeping roosts consisting of a few to several dozen individuals (Young and Thomason 1975; Cook et al. 1976).
In a study of the Zebra Longwing in Costa Rica, Cook et al. (1976) observed a maximum lifespan of 133 days, but estimated the typical adult lifespan to be around 40-50 days, with evidence of senescence (declining survival rate) after about 25 days.
In a study in south Florida, Fleming et al. (2005) estimated the maximum adult lifespan to be 13-14 weeks (91-98 days). Most adults, however, lived less than one month, so average lifespan was estimated to be less than eight weeks (<56 days).
Females lay eggs on young host leaves at branch tips, depositing just a few eggs each day over a period of several months (Opler and Krizek 1984).
Molecular Biology and Genetics
Statistics of barcoding coverage: Heliconius charithonia
Public Records: 18
Specimens with Barcodes: 45
Species With Barcodes: 1
Barcode data: Heliconius charithonia
There are 14 barcode sequences available from BOLD and GenBank. Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species. See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
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Download FASTA File
National NatureServe Conservation Status
Rounded National Status Rank: N5 - Secure
NatureServe Conservation Status
Rounded Global Status Rank: G5 - Secure
Reasons: Widespread and common in neotropics, apparently secure in southern Florida.
Degree of Threat: D : Unthreatened throughout its range, communities may be threatened in minor portions of the range or degree of variation falls within natural variation
Comments: Tolerates disturbance short of complete clearing.
Global Protection: Unknown whether any occurrences are appropriately protected and managed
The caterpillars are white with black spots and have numerous black spikes along their body. Adult butterflies are monomorphic of medium size with long wings. On the dorsal side, the wings are black with narrow white and yellow stripes, with a similar pattern on the ventral side, but paler and with red spots. The wingspan ranges from 72 to 100 mm.
Adults roost in groups of up to 60 individuals on a nightly basis, returning to the same roost every night. These roosts provide protection to adults, the large groups deterring predators and retaining warmth.
The caterpillar feeds on Yellow Passionflower (Passiflora lutea), Corky-stemmed Passionflower (Passiflora suberosa), and Two-flower Passionflower (Passiflora biflora). The adults are unusual among butterflies in that they eat pollen as well as sip nectar. This ability contributes to their longevity—the Zebra Longwing can live up to 6 months as an adult.
Adults exhibit pupal mating in which males wait for a female to emerge from her pupa. Upon emergence, two or more males may fight in order to win a copulation. The winner mates with the females and prevents other males from doing so through a chemical transfer.
During mating, the male passes a nutrient-rich spermatophore to the female that, in addition to providing her valuable proteins, is also thought to reduce the attractiveness of the female to other male mates.
- 1 Distribution
- 2 Roosts
- 3 Feeding
- 4 Mating System
- 5 See also
- 6 Gallery
- 7 References
- 8 External links
It is found in South America, Central America, the West Indies, Mexico, south Texas and peninsular Florida. Adults sometimes migrate north to New Mexico, South Carolina, and Nebraska during the warmer months. It was declared the official butterfly for the state of Florida in the United States in 1996.
It was theorized that H. charithonia did not migrate or disperse over a wide territorial range due to its home range behavior, suggesting that the butterfly stayed near its natal area its entire life, displaying high fidelity to its nightly roost and using specific foraging routes, thereby returning daily to specific sites of adult host plants to collect nectar and pollen and larval plants to mate and lay eggs. Although the butterfly does display some hone range behavior, due to the lack of genetic variation over a wide range of territory, it has been determined that the amount of migration and dispersal in the butterflies was highly underestimated. Therefore, the butterflies do in fact move between territories.
Butterflies with Mexican origins migrate north into Texas due to temperature changes, following the retracting temperature gradient. Rainfall has no effect on migration patterns. Arrival dates and duration of stay at certain locations depends on the distance travelled (the longer the distance travelled, the shorter the duration of stay).
Roosts are formed to avoid attacks from predators since it has been found that there are higher numbers of predatory attacks on solitary individuals versus on those individuals that roost. This follows the prey dilution effect, which states that the probability of an attack on an individual decreases with increasing numbers of individuals in a group. Solitary individuals, or very small roosts, avoid exhibiting proper warning signals so as not to attract predators. On the other hand, too large of groups are overly conspicuous to a predator.
Pre-roosting interactions, which consist of sitting near one another, chasing each other briefly while fluttering, or basking, occur between butterflies from separate roosts, indicating that the butterflies are aware of other roosts in their home range. Despite this, the Zebra Longwing butterflies choose to form smaller aggregations, despite the availability of a substrate to create larger ones. In fact, the optimal or minimum roost size in which predator deterrence is most effective is 5 individuals. However, roost size is also influenced by resource availability and foraging. Aposematism is when prey warn predators of their unprofitability, such as toxicity or unpalatibility, via coloration or patterning. H. charithonia roosts in order to display collective aposematism, thereby deterring predators by displaying their unpalatable taste in a more conspicuous way.
H. charithonia adults form communal roosts on a nightly basis. Communal roosting occurs when individuals aggregate at a particular site for more than a few hours. H. charithonia typically begin roosting as early as 3 hours before sunset and usually leave within two hours after sunrise. Due to the timing of the communal roosting, H. charithonia need to be able to see at low light levels because it helps them navigate when searching for roost sites, either when looking for twigs, tendrils, and dry leaves to land on in order to start a roost or when searching for conspecifics that are already roosting. Their eyes also help them to recognize color patterns in conspecifics. UV rhodopsins in the eye help them to distinguish between 3-OHK yellow pigments, or ultraviolet colors, and other yellow pigments, which to the human eye is undistinguishable. At shorter distances, the butterflies recognize conspecifics via chemical cues. These chemical cues include volatile and nonvolatile substances. The significance of this chemical communication remains largely unknown for Heliconius in general. However, in H. melpomene, (E)-?-ocimene was found to attract males and females in diurnal situations.
It is thought that the behavior of pollen-feeding facilitated the evolution of aposematism and mimicry. Those butterflies that feed on pollen are more distasteful to predators, more brightly colored, and show superior mimetic diversity to those that do not feed on pollen.
Adult butterflies choose their home ranges based on collections of pollen plants. They collect pollen by inserting their proboscis into the flower while making particular movements so that there is secure adhesion to the pollen grains. Digestion occurs immediately after ingestion when the pollen makes contact with saliva and amino acids are dissolved. Optimal amino acid intake occurs via abundant saliva production and gentle and slow mastication. During the night, the butterflies digest pollen since optimal nutritional resources are obtained while resting or sleeping.
Pollen feeding is correlated with higher overall fitness. Individuals who exhibit pollen feeding have a longer lifespan than those who feed only on nectar or sugar water. Females carry more pollen than males since pollen supplies the nutrients necessary for egg production. Amino acids essential for egg development transfer from the pollen to the butterfly eggs. Therefore, oogenesis is greatly affected by pollen intake. When pollen is absent in the diet, oviposition rates decrease and lifetime fecundity, or the number of eggs produced, drops significantly.
Pollen feeding also correlates with the unpalitibility of the butterflies to predators. The amino acids collected from pollen are used as precursors to synthesize cyanogenic glycosides that can be stored in larval and adult tissues, accounting for their toxicity. When pollen availability is low, adults are able to scrounge for nitrogen reserves where there are stored cyanogens. Furthermore, the butterflies are able to recycle the cyanogenic glycosides that they have synthesized previously. With less expectation of pollen quality, females will reallocate their cyanogensto reproductive input since larvae have been seen to benefit the most from cyanogenesis and a lack of amino acids in adult diet does not necessarily correlate with reduced cyanogenic defense.
Larvae regulate their nutritional input to an equal protein-carbohydrate ratio. They feed on the Passiflora plants on which their mother laid their eggs. Passiflora plants have trichomes, which are meant to reduce herbivore attack through physical and/or chemical means. H. charithonia larvae have developed capabilities in order to avoid the effects of the trichomes. They are able to physically free themselves from the entrapment of a trichome by pulling their legs from the hold of the trichome hook and they lay silk mats on the trichomes, providing a surface so that leaf anatomy can be navigated more easily, and they remove the tips of the trichomes by biting them. Therefore, trichome tips are found in the faeces of these individuals. Furthermore, larvae often try to avoid areas where trichome density is highest by staying on the under surface of the leaves.
Male butterflies exhibit a preference for visual, olfactory, tactile, and auditory cues during mating so that females are made more obvious. In H. charithonia, certain host plants provide these cues to males, thereby influencing the time and location of reproduction. This happens because as larvae damage the plant upon eating it, green-leaf volatiles, six carbon alcohols, aldehydes, and acetates, are released. They give olfactory cues to the male, thereby indicating the location of the pupae (mate). Since these pupae have camouflaged coloring and lack strong sexual pheromones, the olfactory cue given off from the damaged plant is necessary for the male to find a mate. Furthermore, the odors given off from the plant not only signal the location of pupae but also provide a trigger that induces learning the location of the plant for future copulations. This is possible due to the fact that the butterflies have a sophisticated spatial memory, as exhibited by their tendency to have specific nocturnal roosting locations and regular visitations of sites with abundant adult resources.
A common problem amongst all butterflies is the ability of mates to discriminate mating with conspecifics and heterospecifics (other butterfly species). Although mistakes do occur, they are quite uncommon and males are able to distinguish, more often than not, between the emissions produced upon the larvae and other herbivores eating the plant. It has been shown that the zebra longwing also uses larval coloration and odors to find mates. Chemical analysis has demonstrated that the larvae release volatiles that are similar chemically to those emitted by the plant.
H. chartihonia also demonstrates pleiotropic mating cues, particularly in regards to mimicry. Pleiotropic mating cues have been simultaneously ecologically and sexually selected. Mimetic patterns have been used in cues to mate-finding as well as in predatory protection, leading to the coevolution of mate choice and assortative mating, a nonrandom mating pattern in which individuals choose mates with similar genotypes and/or phenotypes to themselves. Polymorphism exists in mimetic patterns due to the fact that each morph mimics a different model.
Pupal mating arose once during the evolution of Heliconius, and these species form a clade on the evolutionary tree. Although pupal mating is observed quite frequently in insectaries, it is rarely seen in nature. Males perform precopulatory mate guarding behavior, in which males find and perch on pupae, followed by copulation with the female.
Upon reaching the pupae, males often have to compete to copulate with the female, who is teneral (freshly emerged). Typically, a male visits the same pupa for at least a week, during which time he periodically swarms it, fighting with other males over positioning. Fights consist of males fending off other males that attempt to land on the same pupa by opening their wings. If this does not work, the male tries to throw the intruder off with the pressure of his head and antennae. If more males attempt to swarm the pupa, the two original males will work together to attempt to fend off the others by simultaneously opening their wings, momentarily forgetting that they were originally competitors. Fights usually last one or two hours, but continue throughout the pupa’s development.
The act of pupal mating consists of the male inserting his abdomen into the pupa. If a second male appears, he will fend off other males for the first male by opening his wings while the first male copulates, rather than attempting to mate with the female himself by inserting his abdomen. After two or three hours of mating, the female comes out during which copulation stills occurs for another hour. During the process, females remain relatively still, with the exception of the spreading of their wings and the discharge of meconium. As copulation proceeds, fewer males attempt to approach the female. However, if this does occur, the copulating male continues to fend them off by opening his wings. After copulation is done, the male and female sit side-by-side for some time. During this brief period, no other males attempt to mate with the female.
Nuptial gifts in the form of the spermatophore
Males transfer a protein-rich spermatophore to females upon mating. Spermatophores are thought to be nuptial gifts, which serve a number of different functions. One of these is to provide chemicals (cyanogens) that protect the mother and future offspring from predators so that female and egg survival are enhanced. For females, this is beneficial because egg-laying causes a depletion of protein and her defensive chemicals. Among nine Heliconius species studied, Heliconius chartihonia had the highest average cyanide concentration in its spermatophores.
In most species of butterflies, pheromones play a role in courtship and mate recognition. However, they can also play a role in deterring mates. Spermatophores are also thought to contain anaphrodisiacs, which are pheromones that reduce the attractiveness of the females to subsequent males, thereby indicating that anaphrodisiac evolution is driven by intrasexual selection. Because of this, they help reduce the occurrence of male harassment by already mated females. Furthermore, nonfertile sperm (apyrene) is added to spermatophores to increase the time of refractory periods. Overall, the transfer of anaphrodisiacs is a mechanism for degrading female mating choice.
It has been observed that complete spermatophore degradation to an orange or yellow substance occurs in a 2-week period. Pupal-mating butterflies like Heliconius charatonia are thought to be monandrous, but spermatophore degradation could undermine this idea since spermatophores are often examined to count the number of matings a female has performed. However, upon further observation, it was found that females rarely participate in more than one mating per lifetime.
Sex ratio and distribution
At eclosion, the ratio is highly female-biased, but the rest of the year the sex ratio is overall male-biased (68% males). This is due to the fact that males typically stay near their natal sites to find a mate while females move around to find oviposition or feeding sites at various Passiflora plants. Because females are very mobile, males do not mate with relatives very often, and therefore inbreeding rates are very low.
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- "Attributes of Heliconius charithonia". Retrieved November 14, 2013.
- "Zebra Longwing". Retrieved November 14, 2013.
- "Zebra Heliconian-Florida's State Butterfly!". Retrieved November 14, 2013.
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- Sacledo, Christian. "Behavioral Traits Expressed During Heliconius Butterflies Roost-Assembly". Trop. Lepid. Res 21.2 (2011): 80-83.
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- Bybee, Seth M.; Monica D. Furong Yuan, Jorge Llorente-Bousquets, Robert D. Reed, Daniel Osorio, Adriana D. Briscoe (2012). "UV Photoreceptors and UV-Yellow Wing Pigments in Heliconius Butterflies Allow a Color Signal to Serve Both Mimicry and Intraspecific Communication". The American Naturalist. 1 179: 38–51.
- Sacledo, Christian. The Biology of Heliconius Night Roosting A Foundation. Thesis. UFDC, 2010. Gainesville, Fl: University of Florida, 2010. Print.
- Estrada, Catalina, and Chris D. Jiggins. "Patterns of Pollen Feeding and Habitat Preference among Heliconius Species." Ecological Entomology 27 (2002): 448-56.
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- Cardoso, Márcio Z. "Ecology, Behavior and Binomics: Herbivore Handling of a Plant's Trichome: The Case of Heliconius Charithonia (L.) (Lepidoptera:Nymphalidae) and Passiflora Lobata (Kilip) Hutch. (Passifloraceae)." Neotropical Entomology 37.3 (2008): 247-52.
- Douglas, Matthew M. (1986). The Lives of Butterflies. Ann Arbor: University of Michigan.
- Estrada, Catalina; Lawrence E. Gilbert (2010). "Host Plants and Immatures as Mate-searching Cues in Heliconius Butterflies". Animal Behaviour 80: 231–239.
- Boggs, Carol L., Ward B. Watt, and Paul R. Ehrlich. (2003). Butterflies: Ecology and Evolution Taking Flight. Chicago: University of Chicago.
- Jiggins, Chris D., Igor Emelianov, and James Mallet. (2005). Assortative Mating and Speciation as Pleiotropic Effects of Ecological Adaptation: Examples in Moths and Butterflies. Insect Evolutionary Ecology. By Mark Fellowes, G. J. Holloway, and J. Roff. Wallingford, Oxfordshire: CABI Pub., 451-73.
- Scoble, M. J. (1995). The Lepidoptera: Form, Function and Diversity. [London]: Natural History Museum
- Sourakov, Andrei. (2008). Pupal Mating in Zebra Longwing (Heliconius Charithonia): Photographic Evidence. News of the Lepidopterists' Society 50(1):26-32.
- Cardoso, Márcio Zikán, and Lawrence E. Gilbert. (2006). A Male Gift to Its Partner? Cyanogenic Glycosides in the Spermatophore of Longwing Butterflies (Heliconius). Naturwissenschaften 94(1):39-42.
- Walters, James R., Christine Stafford, Thomas J. Hardcastle, and Chris D. Jiggins. (2012). Evaluating Female Remating Rates in Light of Spermatophore Degradation in Heliconius Butterflies: Pupal-mating Monandry versus Adult-mating Polyandry. Ecological Entomology 37:257-68.
- Fleming, Theodore H., David Serrano, and Jafet Nassar. (2005). Dynamics Of A Subtropical Population Of The Zebra Longwing Butterfly Heliconius Charithonia (Nymphalidae). Florida Entomologist 88(2):169-79.
Names and Taxonomy
Comments: Spelling corrected to original orthography following Opler and Warren (2002). Several spelling variations exist for this species.
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